The Wilson Journal of Ornithology 126(1):47–52, 2014
RELATIVE ABUNDANCE OF SMALL MAMMALS IN NEST CORE
AREAS AND BURNED WINTERING AREAS OF MEXICAN SPOTTED
OWLS IN THE SACRAMENTO MOUNTAINS, NEW MEXICO
JOSEPH L. GANEY,1,6 SEAN C. KYLE,1,2 TODD A. RAWLINSON,1,3
DARRELL L. APPRILL,1,4 AND JAMES P. WARD JR.1,5
ABSTRACT.—Mexican Spotted Owls (Strix occidentalis lucida) are common in older forests within their range but also
persist in many areas burned by wildfire and may selectively forage in these areas. One hypothesis explaining this pattern
postulates that prey abundance increases in burned areas following wildfire. We observed movement to wintering areas
within areas burned by wildfire by four radio-marked Mexican Spotted Owls in the Sacramento Mountains, New Mexico.
These movements occurred during the winters of 2004–2005 and 2005–2006, with some owls migrating in both winters and
others in only one. Wintering areas of these owls occurred within the perimeters of two wildfires that burned in May 2000
and April 2002, respectively. We estimated relative prey abundance and biomass during December 2006 within paired
burned wintering areas and nest core areas used by these owls. Species richness and relative abundance of small mammals
were greater in the burned wintering areas than in the associated nest core areas for all four owls, and estimated prey
biomass ranged from 2–6 times greater in burned wintering areas relative to the paired nest core areas. Burned wintering
areas used by these owls were similar in elevation to their nest core areas, and likely experienced similar weather conditions
during winter. These results suggest that wintering owls moved to areas with greater food resources, rather than to areas
with milder weather. They further suggest that relative prey abundance was greater in burned wintering areas than in the
nest core areas .5 years post-fire, and that these burned wintering areas provided important habitat for Mexican Spotted
Owls in our study area during an energetically stressful season. Received 29 July 2013. Accepted 2 November 2013.
Key words: Mexican woodrat, migration, prey abundance, prey biomass, species richness, voles, wildfire effects.
occupied by Spotted Owls. As a result, wildfire
has now supplanted timber harvest as the greatest
perceived threat to the owl and its habitat (U.S.
Department of the Interior 2012). Thus, understanding the impact of wildfires on population
dynamics and habitat use of Mexican Spotted
Owls is an area of growing interest (U.S.
Department of the Interior 2012).
Wildfire can alter structure in or eliminate the
older forests used by Mexican Spotted Owls over
large areas, leading many observers to assume
that wildfires routinely result in complete loss of
habitat for Mexican Spotted Owls (e.g., Sheppard
and Farnsworth 1995). However, available evidence suggests that these owls often persist in
burned areas, at least in the short term (Bond et al.
2002; Jenness et al. 2004). Similar evidence
exists for the California Spotted Owl (S. o.
occidentalis; Bond et al. 2002, 2009, 2010, 2013;
Roberts et al. 2011; Lee et al. 2012), and
California Spotted Owls have been observed to
selectively forage in burned areas (Bond et al.
2009).
One potential explanation for the continued use
of burned areas by Mexican Spotted Owls
postulates that abundance of the small mammals
that dominate their diet (Ganey 1992, Ward and
Block 1995, Seamans and Gutiérrez 1999, Block
The Mexican Spotted Owl (Strix occidentalis
lucida) occurs in canyonlands and forested
mountains throughout the southwestern United
States and the Republic of Mexico (Gutiérrez
et al. 1995, Ward et al. 1995, U.S. Department of
the Interior 2012). It frequently occupies older
forests or forests with late-seral characteristics
(Ganey and Dick 1995, U.S. Department of the
Interior 2012), and was listed as Threatened under
the Endangered Species Act in 1993, primarily
because of concerns over the loss of older forest
habitat to timber harvest (U.S. Department of the
Interior 1993). Since that time, a number of large
wildfires have burned within the range of the
Mexican Spotted Owl, and many of these fires
have impacted areas that were known to be
1
U.S. Forest Service, Rocky Mountain Research Station,
2500 S. Pine Knoll Drive, Flagstaff, AZ 86001, USA.
2
Current address: Texas Parks and Wildlife Department,
1702 Landmark Lane, Suite 1, Lubbock, TX 79415, USA.
3
Current address: U.S. Forest Service, Lincoln National
Forest, 901 Mechem Road, Ruidoso, NM 88345, USA.
4
Current address: U.S. Forest Service, Lincoln National
Forest, 4 Lost Lodge Road, Cloudcroft, NM 88317, USA.
5
Current address: U.S. Fish and Wildlife Service,
National Wildlife Refuge System, Inventory and Monitoring Branch, Fort Collins, CO 80525, USA.
6
Corresponding author; e-mail: jganey@fs.fed.us
47
48
THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 126, No. 1, March 2014
et al. 2005) may increase following fire, at least in
the short term, and that owls thus may find
favorable foraging habitat in burned areas (Bond
et al. 2002, 2009, 2010; US Department of the
Interior 2012). Studies within the range of
Mexican Spotted Owls suggested increases in
some species of small mammals following
wildfire (Kyle and Block 2000, Converse et al.
2006), but data on this subject generally are sparse
and short term. Thus, it is unknown whether or not
increased prey abundance following fire is a
general pattern, or how long such increases persist
if they do occur.
In conjunction with a study on demography of
Mexican Spotted Owls in the Sacramento Mountains, New Mexico, we observed movement by
four radio-marked owls during winter from nest
core areas in closed-canopy mixed-conifer forests
to areas burned in two large wildfires. Because
winter is suspected to be a time of food shortage
for Mexican Spotted Owls (Block et al. 2005,
Ganey et al. 2005) and prey abundance is
hypothesized to increase post-fire, these moves
might indicate a shift to areas of richer food
resources. We examined species richness, relative
abundance, and biomass of prey at potential
foraging areas around winter roost sites for owls
within the burned wintering areas and at paired
nest core areas used by these owls. Thus, we
provide data on potential differences between
burned wintering areas and nesting areas in prey
abundance and biomass during an energetically
stressful season for Spotted Owls.
STUDY AREA
Our study area was approximately 50,000 ha
in the Sacramento Mountains, south-central New
Mexico, USA. This area encompassed much of
the central portion of the Sacramento Ranger
District, Lincoln National Forest, including the
village of Cloudcroft, New Mexico. Elevation
ranged from 2,000–2,800 m. Terrain consisted of
heavily forested montane slopes and minor
tributaries, with interspersed meadows in the
larger valley bottoms. The predominant forest type
was mixed-conifer, singularly or co-dominated by
white fir (Abies concolor) and Douglas-fir (Pseudotsuga menziesii). Other common tree species
included southwestern white pine (Pinus strobiformis), ponderosa pine (P. ponderosa), and
quaking aspen (Populus tremuloides) (Kaufmann
et al. 1998, Ward 2001). Precipitation averaged
65 cm/yr at Cloudcroft (2,652 m) with summer
thunderstorms providing more than 60% of annual
precipitation and most of the remainder occurring
as winter snowfall (Kaufmann et al. 1998).
Two large wildfires burned within this study
area in 2000 and 2002. The Scott Able fire ignited
on 11 May 2000, and burned 6,213 ha, with
approximately 16 and 25% of this area burned at
moderate and high severity, respectively (Monitoring Trends in Burn Severity 2013). The
Peñasco fire ignited on 30 April, 2002, and
burned 6,073 ha, with approximately 29 and
17% of this area burned at moderate and high
severity, respectively (Monitoring Trends in Burn
Severity 2013).
METHODS
We located general areas occupied by Mexican
Spotted Owls within the study area using
nocturnal calling surveys (Forsman 1983). We
captured owls during daytime follow-up surveys,
using snare poles and baited mist nets, and
attached radio transmitters using a backpack
harness constructed of Teflon ribbon. All owls
discussed here were captured during the summers
of 2004 or 2005, and were radio-tracked though
fall 2006. Radio-marked owls were not located
frequently during the winter months because of
constraints in funding and personnel availability,
as well as access issues. Most locations recorded
for Mexican Spotted Owls were from visual
observations obtained during the day. Consequently, we defined movement to a wintering area
as occurring when a radio-marked owl was
located roosting during the day in an area .2 km
from their nest/roost core (after Ganey and Block
2005) on multiple occasions between 1 November–15 April, with the roost locations separated by
,300 m and occurring over a period of .18 days.
Five radio-marked owls satisfied these criteria,
with four of these owls using areas burned by
wildfires. We established two trapping grids for
each of these four radio-marked owls. One grid
was established within their nest core area (see
below) and the second within their burned
wintering area. We used the nest core area for
comparative purposes, because these areas represented activity centers used during the breeding
season and available foraging habitat which these
owls, by moving to geographically distinct
wintering areas, were not selecting. All nest core
and burned wintering areas were located within
the same forest cover type (mixed-conifer forest).
49
Ganey et al. N MEXICAN SPOTTED OWL PREY IN BURNED WINTERING AREAS
TABLE 1. Summary of migratory movements observed during the winter by five radio-marked Mexican Spotted Owls
in the Sacramento Mountains, New Mexico, 2004–2006.
Owl territory
Owl sex
Fire area used
Winters in which owl
migrated to winter area
Years post-fire
Distance from nest
to winter area (km)
Elevation
difference (m)a
010
027
027
104
067
M
Fb
Mb
F
F
Scott Able
Scott Able
Peñasco
Scott Able
NAc
2004–2005
2004–2005
2004–2005, 2005–2006
2004–2005, 2005–2006
2005–2006
4+
4+
2–3+
4–5+
NA
2.1
12.0
7.1
14.0
9.8
0
0
180
35
600
a
Elevation difference computed as nest elevation – burned wintering area elevation.
Burned wintering areas used by these mated owls were separated from each other by 8.3 km.
This wintering area was not included within a fire perimeter. We considered this an example of altitudinal migration, and did not include nest core or wintering
areas of this owl in small mammal trapping operations.
b
c
Nest core areas (40-ha each) were defined
following Ward and Salas (2000). Wintering areas
were defined by placing a 200-m buffer around all
observed roost locations for each owl using
Arcview 3.2 (ESRI Inc. 1999) then merging these
buffers to create a single unique polygon for each
owl. The 200-m buffers used were arbitrary, but
were intended to define an area close to the
observed roost sites and therefore readily available for foraging owls. We observed these owls
foraging near their roosts during the day on
several occasions.
Trapping grids were established at a randomly
generated location in both nest core and burned
wintering areas. At each grid location, we set 20
traps in two parallel lines (10 traps/line), with 20m spacing between traps. We placed 1 extra-large
(10 3 18 3 60 cm) Sherman live trap at each trap
station. We baited traps with a mixture of peanut
butter, rolled oats, and bird seed, supplied
abundant cotton batting in traps to provide
bedding material for captured animals during cold
winter nights, and covered traps with 1–3 cedar
shingles to provide additional insulation.
We trapped all grids over a 2-week period from
4–15 December 2006. We checked traps in each
grid each morning for 4 days (n 5 80 potential
trap nights/grid). We noted any cases where traps
were sprung without capturing an animal and
subtracted such traps from available trap occasions. We marked each animal captured with a
single numbered ear tag (Monel-1005s1, National
Band and Tag, Newport, Kentucky), to allow
identification of new versus recaptured individuals, and recorded species (or genus for whitefooted mice [Peromyscus spp.]) of the captured
animal. All captured animals were released
unharmed at the point of capture.
For each transect, we report the number of
unique individuals captured by species and total
biomass of all species. We used the number of
unique individuals captured as an index of
abundance, because this index often outperforms
model-based estimators of abundance when data
are sparse (McKelvey and Pearson 2001). Biomass (g) was calculated as the summed biomass
of all unique individuals captured. We used values
presented in Ward (2001; Table 2.8) to generate
mass estimates for individual prey species. Ward
(2001; Table 2.8) presented means for small
mammals trapped in each of four cover types in
the Sacramento Mountains. We averaged these
mean values across the four cover types to
generate our mass estimates. Because capture
data were sparse, we did not conduct formal
statistical analyses and simply present summary
information here.
RESULTS
Five radio-marked owls from four unique owl
territories moved from their breeding area to
geographically distinct wintering areas, based on
our movement criteria (Table 1). One owl moved
to a wintering area 600 m lower in elevation than
the nest core area and located outside of the fire
perimeters. We considered this an example of
altitudinal migration (Ganey and Block 2005) and
did not trap small mammals in this wintering area.
Only two species of small mammals (whitefooted mice and Mexican woodrats [Neotoma
mexicana]) were captured in nest core areas, with
white-footed mice accounting for 85% of individuals captured (Table 2). In contrast, four
species were captured in burned wintering areas
(white-footed mice [72% of individuals captured],
Mexican woodrats, and both long-tailed [Microtus
50
THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 126, No. 1, March 2014
TABLE 2. Numbers of small mammals captured by species and estimated biomass of small mammals trapped in nest
core and burned wintering areas used by four radio-marked Mexican Spotted Owls in the Sacramento Mountains, New
Mexico, 2004–2006.
Number of unique individuals captured
Territory
Owl sex
Grid type
Trap occasionsa
White-footed
mouse
010
M
027
Fc
027
Mc
104
F
Nest core
Winter area
Nest core
Winter area
Nest core
Winter area
Nest core
Winter area
78
74
76
75
76
72
73
77
4
4
5
18
5
4
2
11
Mexican
woodrat
Long-tailed
vole
Mogollon
vole
Biomass (g)b
2
4
0
1
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
3
0
0
0
0
306.8
577.9
86.0
507.2
86.0
187.8
34.4
189.2
a
Calculated as (number of traps 34 occasions) minus number of sprung or otherwise unavailable traps.
Biomass estimates used were derived by averaging estimates from Ward (2001: table 2.8) across four cover types. Estimates used here were: white-footed mouse
5 17.2 g, Mexican woodrat 5 119.0 g, long-tailed vole 5 33.1 g, and Mogollon vole 5 26.2 g.
c
The same nest core area was used for both the male and female from territory 027.
b
longicaudus] and Mogollon [M. mogollonensis]
voles; Table 2).
Relative prey abundance in winter was low in
all areas, but was greater in burned wintering
areas than in the paired nest core areas (Table 2).
In addition, estimated prey biomass ranged from
approximately double to almost six times greater
in burned wintering areas relative to the paired
nest core areas. Total biomass of individuals
captured in burned wintering areas was 237.2 g
greater than biomass captured in nest core areas,
on average (95% CI 5 12.0–462.5 g), despite
having more trap occasions in the nest core area in
three of four cases (Table 2).
DISCUSSION
Our results support the hypothesis that prey
resources were greater in the burned wintering
areas than in the paired nest core areas. Species
richness, relative abundance, and biomass of
small mammals during the trapping period all
were greater within the burned wintering areas
than within the paired nest cores of the radiomarked owls. Because the Scott Able fire burned
in May 2000 and we trapped small mammals in
that area during December 2006, our results
suggest that prey abundance in the burned
wintering areas was greater than in the nest core
areas even where those burned wintering areas
were .6 years post-fire.
Although many radio-marked Mexican Spotted
Owls in previous studies remained in their
breeding areas during winter, some individuals
in all populations studied migrated to wintering
areas at lower elevations (Ganey and Block 2005:
Table 2). Such movements typically allowed
migrating owls to winter in areas featuring
warmer temperatures below the level of persistent
snow, but also could be driven by relative prey
availability. Only one study (Block et al. 2005),
involving two radio-marked owls from a single
territory, quantified differences in prey abundance
between lower elevation wintering areas and nest
core areas. These owls moved ,40 km from their
nesting area in ponderosa pine – Gambel oak
(Quercus gambelii) forest to a wintering area in
pinyon (Pinus spp.) – juniper (Juniperus spp.)
woodland, ,920 m lower in elevation than the
nest area. Prey biomass during the winter was
approximately seven times greater in the wintering area than in the nesting area of these owls
(Block et al. 2005). Thus, these owls moved a
relatively long distance to a wintering area at
lower elevation in a completely different forest
cover type, which featured both milder weather
and higher prey abundance.
In contrast, the burned wintering areas used by
owls in our study were similar in elevation to the
paired nest core areas, ,15 km from those nest
core areas in all cases, and located in the same
forest cover type. Consequently, these owls did
not appear to be moving to areas with more
favorable weather conditions, such as warmer
temperatures or reduced snow cover. The fact that
they moved to areas with richer food resources
suggests that they may have been seeking greater
prey abundance during a season when prey for
Mexican Spotted Owls are suspected to be scarce
Ganey et al. N MEXICAN SPOTTED OWL PREY IN BURNED WINTERING AREAS
(Ward 2001, Block et al. 2005, Ganey et al. 2005).
Bond et al. (2010) also documented expanded
winter movements and use of burned areas by
radio-marked California Spotted Owls, and Irwin
et al. (2013) documented preferential use of
harvested areas by Northern Spotted Owls (S. o.
caurina) during winter. Irwin et al. (2013)
suggested that such use may have been a response
to greater prey abundance in these areas.
Clearly, our data were sparse, and involved a
limited trapping effort in one time period and in
burned wintering and nest core areas of only four
owls. We were not able to estimate detection
probabilities, and so cannot rule out the possibility
that trappability of small mammals differed
between nest cores and burned wintering areas,
although we have no biological reason to suspect
such a difference. We also were not able to
generate estimates of precision around abundance
estimates, which limits the strength of our
comparisons between paired nest core and burned
wintering areas (McKelvey and Pearson 2001).
The observed differences between paired nest
core and burned wintering areas were both
relatively large and consistent in direction, and
the index of abundance that we used has been
shown to perform well in many situations
(McKelvey and Pearson 2001).
Finally, because our study was opportunistic
and observational rather than experimental, we
cannot conclusively attribute differences in prey
abundance between paired nest core and burned
wintering areas to the effects of wildfire. Paired
nest and burned wintering areas sampled were
separated in all cases by ,15 km, and were
similar in elevation and located in the same forest
cover type. Thus, the primary difference between
these paired areas was that wintering areas were
burned and nest cores were not. Consequently, it
seems likely that fire effects at least partly
explained the observed differences in prey
abundance between these paired sites.
It would be desirable to confirm our results
with additional and more intensive trapping over
longer time frames and on more than one
occasion, in multiple seasons, and in more
geographic areas throughout the range of Mexican
Spotted Owls. In the meantime, however, our
results suggest that the burned wintering areas
used supported greater species richness, relative
abundance, and biomass of prey than the paired
nest core areas, that these differences were
observed as much as .6 yrs post-fire, and that
51
in our study area these burned wintering areas
provided important habitat for Mexican Spotted
Owls during an energetically stressful time of
year.
ACKNOWLEDGMENTS
We thank the many dedicated field personnel who helped
locate, capture, and radio-track owls within the Sacramento
Mountains. We also thank personnel on the Sacramento
Ranger District, Lincoln National Forest (especially M.
Mauter, J. Montoya, R. Guaderrama, D. Salas, and J.
Williams), for operational support during the study. Major
funding was provided by the Southwestern Region, USFS,
with additional funding from the Lincoln National Forest
and Rocky Mountain Research Station, USFS. We thank D.
DeLorenzo for his support and assistance with securing
funding throughout the study, and two anonymous reviewers for helpful comments on an earlier draft of this paper.
LITERATURE CITED
BLOCK, W. M., J. L. GANEY, P. E. SCOTT, AND R. M. KING.
2005. Prey ecology of the Mexican Spotted Owl in
ponderosa pine-Gambel oak forests of northern
Arizona. Journal of Wildlife Management 69:618–629.
BOND, M. L., R. J. GUTIÉRREZ, A. B. FRANKLIN, W. S.
LAHAYE, C. A. MAY, AND M. E. SEAMANS. 2002.
Short-term effects of wildfires on Spotted Owl
survival, site fidelity, and mate fidelity. Wildlife
Society Bulletin 30:1022–1028.
BOND, M. L., D. E. LEE, AND R. B. SIEGEL. 2010. Winter
movements by California Spotted Owls in a burned
landscape. Western Birds 41:174–180.
BOND, M. L., D. E. LEE, R. B. SIEGEL, AND M. W. TINGLEY.
2013. Diet and home-range size of California Spotted
Owls in a burned forest. Western Birds 44:114–126.
BOND, M. L., D. E. LEE, R. B. SIEGEL, AND J. P. WARD JR.
2009. Habitat use and selection by California Spotted
Owls in a postfire landscape. Journal of Wildlife
Management 73:1116–1124.
CONVERSE, S. J., G. C. WHITE, AND W. M. BLOCK. 2006.
Small mammal responses to thinning and wildfire in
ponderosa pine-dominated forests of the southwestern
United States. Journal of Wildlife Management
70:1711–1722.
ESRI INC. 1999. ArcView Version 3.2. Environmental
Systems Research Institute, Redlands, California,
USA.
FORSMAN, E. D. 1983. Methods and materials for locating
and studying Spotted Owls. USDA Forest Service,
General Technical Report PNW-162. Pacific Northwest Research Station, Portland, Oregon, USA.
GANEY, J. L. 1992. Food habits of Mexican Spotted Owls in
Arizona. Wilson Bulletin 104:321–326.
GANEY, J. L. AND W. M. BLOCK. 2005. Winter movements
and range use of radiomarked Mexican Spotted Owls:
An evaluation of current management recommendations. USDA Forest Service, General Technical Report
RMRS-GTR-148-WWW. Rocky Mountain Research
Station, Fort Collins, Colorado, USA.
52
THE WILSON JOURNAL OF ORNITHOLOGY N Vol. 126, No. 1, March 2014
GANEY, J. L., W. M. BLOCK, B. E. STROHMEYER, AND J. P.
WARD JR. 2005. Home range, habitat use, survival, and
fecundity of Mexican Spotted Owls in the Sacramento
Mountains, New Mexico. Southwestern Naturalist
50:323–333.
GANEY, J. L. AND J. L. DICK JR. 1995. Habitat relationships
of Mexican Spotted Owls: current knowledge. Chapter
4:1–42 in Recovery plan for the Mexican Spotted Owl.
U.S. Fish and Wildlife Service. Albuquerque, New
Mexico, USA.
GUTIÉRREZ, R. J., A. B. FRANKLIN, AND W. S. LAHAYE.
1995. Spotted Owl (Strix occidentalis). The birds of
North America, Number 179.
IRWIN, L. L., D. F. ROCK, AND S. C. ROCK. 2013. Do
Northern Spotted Owls use harvested areas? Forest
Ecology and Management 310:1029–1305.
JENNESS, J. S., P. BEIER, AND J. L. GANEY. 2004.
Associations between forest fire and Mexican Spotted
Owls. Forest Science 50:765–772.
KAUFMANN, M. R., L. S. HUCKABY, C. M. REGAN, AND J.
POPP. 1998. Forest reference conditions for ecosystem
management in the Sacramento Mountains, New
Mexico. USDA Forest Service, General Technical
Report RMRS-GTR-19. Rocky Mountain Research
Station, Fort Collins, Colorado, USA.
KYLE, S. C. AND W. M. BLOCK. 2000. Effects of wildfire
severity on small mammals in northern Arizona
ponderosa pine forests. Pp. 163–168 in Fire and forest
ecology: innovative silviculture and vegetation management (W. K. Moser and C. F. Moser, Editors). Tall
Timbers Fire Ecology Conference Proceedings 21.
Tall Timbers Research Station. Tallahassee, Florida,
USA.
LEE, D. E., M. L. BOND, AND R. B. SIEGEL. 2012. Dynamics
of breeding-season site occupancy of the California
Spotted Owl in burned forests. Condor 114:792–802.
MCKELVEY, K. S. AND D. E. PEARSON. 2001. Population
estimation with sparse data: the role of estimators
versus indices revisited. Canadian Journal of Zoology
79:1754–1765.
MONITORING TRENDS IN BURN SEVERITY (MTBS). 2013. MTBS
data. U.S. Geological Survey and U.S. Forest Service.
www.mtbs.gov/index.html (accessed 21 Mar 2013).
ROBERTS, S. L., J. W. VAN WAGTENDONK, A. K. MILES, AND
D. A. KELT. 2011. Effects of fire on Spotted Owl site
occupancy in a late-successional forest. Biological
Conservation 144:610–619.
SEAMANS, M. E. AND R. J. GUTIÉRREZ. 1999. Diet
composition and reproductive success of Mexican
Spotted Owls. Journal of Raptor Research 33:143–148.
SHEPPARD, G. AND A. FARNSWORTH. 1995. Fire effects and
use of prescribed fire in Mexican Spotted Owl habitat.
Pages 131–135 in Proceedings – fire effects on rare
and endangered species and habitats Conference. 13–
16 November 1995, Coeur d’Alene, Idaho. International Association of Wildland Fire.
U.S. DEPARTMENT OF THE INTERIOR. 1993. Endangered and
threatened wildlife and plants: final rule to list the
Mexican Spotted Owls as a threatened species. U.S.
Fish and Wildlife Service. Federal Register 58:14248–
14271.
U.S. DEPARTMENT OF THE INTERIOR. 2012. Final recovery
plan for the Mexican Spotted Owl (Strix occidentalis
lucida), First Revision. U.S. Fish and Wildlife Service.
Albuquerque, New Mexico, USA.
WARD, J. P. JR. 2001. Ecological responses by Mexican
Spotted Owls to environmental variation in the
Sacramento Mountains, New Mexico. Dissertation,
Colorado State University, Fort Collins, USA.
WARD, J. P. JR. AND W. M. BLOCK. 1995. Mexican Spotted
Owl prey ecology. Chapter 5:1–48 in Recovery Plan
for the Mexican Spotted Owl (Strix occidentalis
lucida), Volume II - Technical supporting information.
USDI Fish and Wildlife Service. Albuquerque, New
Mexico, USA.
WARD, J. P. JR., A. B. FRANKLIN, S. E. RINKEVICH, AND F.
CLEMENTE. 1995. Distribution and abundance of
Mexican Spotted Owls. Chapter 1:1–14 in USDI Fish
and Wildlife Service, Recovery Plan for the Mexican
Spotted Owl (Strix occidentalis lucida), Volume 2
Technical supporting information. Albuquerque, New
Mexico, USA.
WARD, J. P. JR. AND D. SALAS. 2000. Adequacy of roost
locations for defining buffers around Mexican Spotted
Owl nests. Wildlife Society Bulletin 28:688–698.